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Morphology and Evolution of Turtles Vertebrate Paleobiology and Paleoanthropology Series Edited by Eric Delson Vertebrate Paleontology, American Museum of Natural History, New York, NY 10024, USA [email protected] Eric J. Sargis Anthropology, Yale University New Haven, CT 06520, USA [email protected] Focaltopicsforvolumesintheserieswillincludesystematicpaleontologyofallvertebrates(fromagnathanstohumans), phylogeny reconstruction, functional morphology, Paleolithic archaeology, taphonomy, geochronology, historical biogeography, and biostratigraphy. Other fields (e.g., paleoclimatology, paleoecology, ancient DNA, total organismal community structure) may be considered if the volume theme emphasizes paleobiology (or archaeology). Fields such as modeling of physical processes, genetic methodology, nonvertebrates or neontology are out of our scope. Volumes in the series may either be monographic treatments (including unpublished but fully revised dissertations) or edited collections, especially those focusing on problem-oriented issues, with multidisciplinary coverage where possible. Editorial Advisory Board Nicholas Conard (University of Tübingen), John G. Fleagle (Stony Brook University), Jean-Jacques Hublin (Max Planck Institute for Evolutionary Anthropology), Ross D.E. MacPhee (American Museum of Natural History), Peter Makovicky(TheFieldMuseum),SallyMcBrearty(UniversityofConnecticut),JinMeng(AmericanMuseumofNatural History), Tom Plummer (Queens College/CUNY), Mary Silcox (University of Toronto). For other titlespublished in thisseries, goto www.springer.com/series/6978 Morphology and Evolution of Turtles Proceedings of the Gaffney Turtle Symposium (2009) in Honor of Eugene S. Gaffney Edited by Donald B. Brinkman Royal Tyrrell Museum, Midland Provincial Park, Highway 838 Drumheller, AB T0J 0Y0, Canada Patricia A. Holroyd Museum of Paleontology, University of California, 1101 Valley Life Sciences Building, Berkeley, CA 94720-4780, USA James D. Gardner Royal Tyrrell Museum, Midland Provincial Park, Highway 838 Drumheller, AB T0J 0Y0, Canada 123 Editors Donald B. Brinkman James D.Gardner Royal TyrrellMuseum Royal TyrrellMuseum Midland Provincial Park Midland Provincial Park Highway 838 Highway 838 Drumheller, ABT0J 0Y0 Drumheller, ABT0J 0Y0 Canada Canada Patricia A.Holroyd Museumof Paleontology Universityof California 1101Valley Life Sciences Building Berkeley,CA 94720-4780 USA ISSN 1877-9077 ISSN 1877-9085 (electronic) ISBN 978-94-007-4308-3 ISBN 978-94-007-4309-0 (eBook) DOI 10.1007/978-94-007-4309-0 Springer Dordrecht Heidelberg New YorkLondon LibraryofCongressControlNumber:2012937464 (cid:2)SpringerScience+BusinessMediaDordrecht2013 Chapter24:FossilEuropeanSeaTurtles:AHistoricalPerspective(cid:2)NHMTheNaturalHistoryMuseum2012 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartofthematerialis concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed.Exemptedfromthislegalreservationarebriefexcerptsinconnectionwithreviewsorscholarlyanalysis ormaterialsuppliedspecificallyforthepurposeofbeingenteredandexecutedonacomputersystem,forexclusive use by the purchaser of the work. Duplication of this publication or parts thereof is permitted only under the provisions of the Copyright Law of the Publisher’s location, in its current version, and permission for use must always be obtained from Springer. Permissions for use may be obtained through RightsLink at the Copyright ClearanceCenter.ViolationsareliabletoprosecutionundertherespectiveCopyrightLaw. Theuseofgeneraldescriptivenames,registerednames,trademarks,servicemarks,etc.inthispublicationdoesnot imply,evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevantprotectivelawsand regulationsandthereforefreeforgeneraluse. ProductLiability:Whiletheadviceandinformationinthisbookarebelievedtobetrueandaccurateatthedateof publication,neithertheauthorsnortheeditorsnorthepublishercanacceptanylegalresponsibilityforanyerrorsor omissions that may be made. The publisher makes no warranty, express orimplied, with respect to the material containedherein. CoverIllustration:‘‘TheGhostsofTurtlesPast’’byDonBrinkman Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) Preface With over 100 publications, many of monographic length, Gene Gaffney has brought researchonfossilturtlestotheforefrontofpaleontologicalstudy.Hisworksetthestage for the current explosion of research on fossil turtles, which is showing exceptional potential for contributing to higher level concepts, such as the nature of evolutionary processes, paleobiogeography, and paleoecology. Upon Gene’s retirement in 2007, the consensusamongturtleresearcherswasthathisvaluablecontributiontoturtleresearch should be recognized. An informal survey of researchers at the 2007 Society of Vertebrate Paleontology meetings indicated that a dedicated meeting on fossil turtles and turtle relationships, together with a Festschrift publication, was the most appro- priate way to honor Gene. The idea of a stand-alone meeting dedicated to fossil turtles follows a tradition establishedbyFranceLapparent,whoorganizedthefirstinternationalmeetingonfossil turtles in Paris, France, in 1983. Subsequent fossil turtle symposia were held at: the American Museum of Natural History, New York, USA, in 1987; the University of AlbertainEdmonton,Alberta,Canada(inconjunctionwiththe75thannualmeetingof the American Society of Ichthyologists and Herpetologists), in 1995; and at the Zoological Institute of the Russian Academy of Sciences in St. Petersburg, Russia, in 2003.Atthetimeofwriting,plansarewellunderwayforhostingthenextsymposiumat the University of Tu¨bingen, Germany, in 2012. To emphasize that the meeting was being held in Gene’s honor, it was formally named the ‘‘Gaffney Turtle Symposium’’, and quickly became known simply as ‘‘Gaffneyfest’’.ThesuccessofthemeetingwasproofofthehighesteeminwhichGeneis valued by his peers. Held at the Royal Tyrrell Museum of Palaeontology in October 2009, the meeting lasted 2 full days, featured 37 talks and 16 posters, and was attended by approximately 100 people. In keeping with the desire to mark Gene’s latest career milestone,themeetingwasagatheringofseveralgenerationsofresearchers.Oneendof thespectrumwasrepresentedby6ofthe13attendeesattheoriginal1984Parismeeting and by 2 of Gene’s classmates from university, all of whom made presentations. The other end of the generational spectrum was represented by an equal number of pre- sentations from researchers who were either within the final stages of their graduate training or had graduated within the past year. And somewhere in the middle were presenters who had never known a time when some of Gene’s classic papers, such as ‘‘ThesystematicsoftheNorthAmericanFamilyBaenidae(Reptilia,Cryptodira)’’(1972) and ‘‘Comparative cranial morphology of Recent and fossil turtles’’ (1979) were not available as standard references. Many of the previous turtle meetings resulted in benchmark publications that have playedakeyroleinthedevelopmentoffossilturtleresearch,andinsomecases,continue to be standard references. The meeting in Paris in 1984 resulted in the first volume of v vi Preface Vertebrata Paleochelonica, a series that included, among other topics, reviews of fossil turtles from central Asia and Mongolia that provided western researchers with the first easilyaccessibleoverviewofturtleresearchbeingundertakenintheSovietUnion.During the second meeting at the American Museum of Natural History in 1987, attendees received a compilation of translated papers on fossil turtles that fostered further devel- opment of an international perspective by turtle researchers. Thus, it was clear in the organizers’mindsthattohavealong-lastingimpact,theGaffneyTurtleSymposiummust befollowedbyaFestschriftvolume.Itwasrecognizedearlyonbytheeditorialteamthat tobeafittingtribute,theFestschriftideallyshouldcontainamixoffocusedpapersthat wouldappealtospecialistsalongwithpapershavingabroaderscopethatwouldappealto amoregeneralaudience.Forthelatter,wesolicitedseveralreview-stylepapersonsubjects of current interest, such as the origins of turtles and development of the turtle shell. Althoughtherearesomeobviousgapsinthisvolume(e.g.,nocomprehensiveanalysesof turtle relationships or paleobiogeography), this Festschrift contains a broad range of researchandreviewpapers,andwehopeitwillbeausefulresourceforyearstocome. Don Brinkman got the ball rolling on this Festschrift project. Soon thereafter, Jim GardnerandPatHolroydsteppeduptohelpthevolumecometofruition,bringingwith them extensive experience as editors for the Journal of Vertebrate Paleontology. Due to theenthusiasticresponsetoourinvitationtocontributetoGene’sFestschrift,theeditors realizedthevolumewouldhavetotaketheformofabook,ratherthanaspecialissueof the Bulletin of the American Museum of Natural History as originally planned. At the suggestionofWalterJoyce,theeditorsapproachedEricDelsonandEricSargiswiththe ideaofincludingthisinSpringer’sVertebratePaleobiologyandPaleoanthropologyseries. PatHolroyddidthekeyworkindraftingtheproposal,whichwaspositivelyreceivedby Springer. Don Brinkman and Jim Gardner shared the workload in getting the manu- scriptsandotherpartsofthetextreadyforpublication.Severalyearslater,thisvolume is the final result of the editors’ and authors’ collective efforts. Fig.1 LogofortheGaffneyTurtleSymposium Fig.2 Traditional Chinese paintingpresented to EugeneGaffney duringthe Gaffney Turtle Symposium in October2009 Acknowledgments Many of the papers included in this volume were presented at the Gaffney Turtle Symposium,heldinOctober2009attheRoyalTyrrellMuseumPalaeontologyinhonor of Gene Gaffney. This symposium was made possible through the financial support of The Cooperating Society of the Royal Tyrrell Museum, the American Museum of Natural History, and a Community Spirit Grant from the Department of Culture and Community Spirit of the Province of Alberta. The support of those organizations is greatly appreciated, especially because it helped defray travel expenses for several international participants who otherwise would not have been able to attend the sym- posium.Thesymposiumalsowouldnothavebeenpossiblewithouttheassistanceofthe organizingcommitteeandstaffoftheRoyalTyrrellMuseum.Aswell,theeditorsthank the participants of the symposium, whose enthusiastic discussions led to the develop- ment and refinement of many of the ideas presented in this volume. Theroleofreviewersinthepeer-reviewprocessiscriticalforavolumeofthiskind.As wellasgenerouslydonatingtheirtimeandenergy,thereviewersforthisvolumemaintained a high degree of professionalism, provided constructive and supportive comments, and returnedtheirreviewspromptly;allofthismadetheworkoftheeditorsmucheasier.For theireffortsinreviewingmanuscripts,theeditorsthankthefollowingindividuals: David Brockhurst, Richard Moody, Ann Burke, Gary Morgan, Edwin Cadina, Jacqueline Moustakas, Bob Carroll, Hiroshi Nagashima, Zolta´n Csiki, Jim Parham, Igor Danilov, Larry Powell, Mike Everhart, Ada´n Pe´rez-Garcı´a, Andrew Farke, Ma´rton Rabi, Nadia Fro¨bisch, Olivier Rieppel, Marcelo de la Fuente, Pedro Seyferth Romano, Emiliano Jime´nez Fuentes, Krister Smith, Gene Gaffney, Juliana Sterli, Donald Henderson, Steven Sweetman, Ren Hirayama, Anthony Russell, Howard Hutchison, Michael Ryan, Walter Joyce, Juliana Sterli, Ben Kear, Haiyan Tong, Takuya Konishi, Matthew Vickaryous, France Lapparent de Broin, Michael Wuttke, Derek Larson, Chang-Fu Zhou, Peter Meylan, and several anonymous reviewers. Andrew Milner, vii viii Acknowledgments Three of the above-listed reviewers (Igor Danilov, Walter Joyce, and Jim Parham) deserve particular recognition, because they were called upon repeatedly and often at short notice to review manuscripts for this volume. We are grateful to Eric Delson and Eric Sargis (a.k.a. ‘‘the Erics’’), co-editors for Springer’s Vertebrate Paleobiology and Paleoanthropology Book Series, for their help and guidance in shepherding this project from proposal to publication. Judith Terpos, TamaraWelschot,andotherstaffatSpringeralsoprovidedinvaluablehelpinproducing this volume. The Cooperating Society of the Royal Tyrrell Museum provided funds to cover additional costs for printing some of the color figures in this volume. Finally, we thank Gene Gaffney for his leadership and willingness to share infor- mation and ideas about fossil turtles, paleontology, and science throughout his more than thirty year career as a student of fossil turtles. His passionate interest in turtle evolutionandtheinformationtheyprovideforstudyingtheprocessesandpatternsthat have shaped the history of life, his openness in sharing unpublished information and ideas, and his willingness to provide access to undescribed specimens have led to the development of a uniquely collegiate atmosphere among fossil turtle researchers. We hope this book, and the symposium that preceded it, demonstrate the esteem and respect with which Gene and his work are held. Drumheller, Alberta, Canada, 13 January 2012 Donald B. Brinkman and James D. Gardner Berkeley, California, USA, 13 January 2012 Patricia A. Holroyd Contents Part I Perspectives on the Life and Accomplishments of Eugene S. Gaffney 1 Eugene S. Gaffney: A Professional Biography and Bibliography. . . . . . . . . 3 Robert L. Carroll 2 Autobiography (Through May 2009). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Eugene S. Gaffney Part II The Origin of Turtles 3 Problems of the Ancestry of Turtles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Robert L. Carroll 4 Origin of the Turtle Body Plan: The Folding Theory to Illustrate Turtle-Specific Developmental Repatterning . . . . . . . . . . . . . . . . . . . . . . . 37 Hiroshi Nagashima, Shigehiro Kuraku, Katsuhisa Uchida, Yoshie Kawashima-Ohya, Yuichi Narita and Shigeru Kuratani 5 The Evolution of the Turtle Shell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Olivier Rieppel 6 Three Ways to Tackle the Turtle: Integrating Fossils, Comparative Embryology, and Microanatomy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 Torsten M. Scheyer, Ingmar Werneburg, Christian Mitgutsch, Massimo Delfino and Marcelo R. Sánchez-Villagra 7 Geometric and Developmental Perspectives on the Evolution of the Skull and Internal Carotid Circulation in Turtles . . . . . . . . . . . . . . 71 Tetsuto Miyashita Part III The Early Diversification of Turtles 8 New Material of the Platychelyid Turtle Notoemys zapatocaensis from the Early Cretaceous of Colombia; Implications for Understanding Pleurodira Evolution . . . . . . . . . . . . . . . . . . . . . . . . . . 105 Edwin A. Cadena, Carlos A. Jaramillo and Jonathan I. Bloch ix x Contents 9 Spoochelys ormondea gen. et sp. nov., an Archaic Meiolaniid-Like Turtle from the Early Cretaceous of Lightning Ridge, Australia. . . . . . . . . . . . . . 121 Elizabeth T. Smith and Benjamin P. Kear 10 Turtles from the Jurassic Shishugou Formation of the Junggar Basin, People’s Republic of China, with Comments on the Basicranial Region of Basal Eucryptodires. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147 Donald B. Brinkman, David A. Eberth, Xing Xu, James M. Clark and Xiao-Chun Wu 11 Rediscovery of the Carapace of the Lost Holotype of the Purbeck Turtle Chelone obovata Owen 1842. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173 Andrew R. Milner 12 Kappachelys okurai gen. et sp. nov., a New Stem Soft-Shelled Turtle from the Early Cretaceous of Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179 Ren Hirayama, Shinji Isaji and Tsuyoshi Hibino 13 Morphology and Relationships of Brachyopsemys tingitana gen. et sp. nov. from the Early Paleocene of Morocco and Recognition of the New Eucryptodiran Turtle Family: Sandownidae . . . . . . . . . . . . . . . . . . . 187 Haiyan Tong and Peter Meylan Part IV Pleurodire Diversity and Biogeography 14 A New Long-Necked Turtle, Laganemys tenerensis (Pleurodira: Araripemydidae), from the Elrhaz Formation (Aptian–Albian) of Niger . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215 Paul C. Sereno and Sara J. ElShafie 15 Two Synchronic and Sympatric Bothremydidae Taxa (Chelonii, Pleurodira) in the Late Cretaceous Site of ‘‘Lo Hueco’’ (Cuenca, Spain). . . . . . . . . . . . 251 Adán Pérez-García, Francisco Ortega and Xabier Murelaga 16 New Information about Pelomedusoides (Testudines: Pleurodira) from the Cretaceous of Brazil. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261 Pedro S. R. Romano, Gustavo R. Oliveira, Sergio A. K. Azevedo, Alexander W. A. Kellner and Diogenes de Almeida Campos 17 Nostimochelone lampra gen. et sp. nov., an Enigmatic New Podocnemidoidean Turtle from the Early Miocene of Northern Greece . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277 Georgios L. Georgalis, Evangelos Velitzelos, Dimitrios E. Velitzelos and Benjamin P. Kear 18 A New Species of Bairdemys (Pelomedusoides: Podocnemididae) from the Oligocene (Early Chattian) Chandler Bridge Formation of South Carolina, USA, and Its Paleobiogeographic Implications for the Genus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289 Robert E. Weems and James L. Knight

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